3990 (4): 575 583 Article · Archipelago. Similar information is repeated in FishBase (Froese &...

Accepted by R. Pethiyagoda: 7 Jul. 2015; published: 28 Jul. 2015

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ZOOTAXA

ISSN 1175-5326 (print edition)

ISSN 1175-5334 (online edition)Copyright © 2015 Magnolia Press

Zootaxa 3990 (4): 575–583

www.mapress.com/zootaxa/

Article

575

http://dx.doi.org/10.11646/zootaxa.3990.4.6

http://zoobank.org/urn:lsid:zoobank.org:pub:7FA710EF-3947-4E7F-AAAF-D6BAC7FD72BA

Why is Pseudosphromenus cupanus (Teleostei: Osphronemidae) reported from

Bangladesh, Indonesia, Malaysia, Myanmar, and Pakistan?

SVEN O. KULLANDER1

, MD. MIZANUR RAHMAN2

, MICHAEL NORÉN1

& ABDUR ROB MOLLAH2

1

Department of Zoology, Swedish Museum of Natural History, PO Box 50007, SE-104 05 Stockholm, Sweden.

E-mail: [email protected]; [email protected]

2

Department of Zoology, University of Dhaka , Dhaka-1000, Bangladesh.

E-mail: [email protected]; [email protected]

Abstract

The native distribution of the small labyrinth fish species Pseudosphromenus cupanus includes southern India and Sri

Lanka. According to literature it has a range including also Pakistan, Bangladesh, Myanmar, Malaysia, and Indonesia (Su-

matra) but there are no voucher specimens or reliable observations from those areas. The distribution record of P. cupanus

was inflated partly by including P. dayi as a synonym. Pseudosphronemus dayi is native to the Western Ghats in India, but

the origin of the aquarium importation in 1907 was reported as both Cochin (=Kochi) and Malacca (=Malaysia), the latter

locality obviously in error. The basis for the Sumatra record is an obviously mislabeled sample of P. dayi from Pulau Weh

close to Sumatra. The basis for reporting the species from Pakistan, Myanmar or Bangladesh could not be located. Mis-

identified museum specimens from Myanmar and Pakistan identified as P. cupanus were never published on. Pseudosph-

romenus cupanus has been considered recently to be extinct in Bangladesh, but in fact it never occurred there.

Key words: Asia, Freshwater, Geographical distribution, Threat status

Introduction

Pseudosphromenus cupanus (Cuvier, 1831) is a small species, about 60 mm long in total length, of the labyrinth-

fish family Osphronemidae (Fig. 1a). It is characterised by its elongate body, relatively long but low dorsal and anal

fins, and the caudal fin ending in a point. It is nearly uniform in colour or with a faint dark band along the side, and

with a black spot at the base of the caudal fin. The only other species in the genus Pseudosphromenus Bleeker,

1879, is P. dayi (Engmann, 1909) which has the same shape and similar dorsal and anal fins, but the caudal fin is

longer (Figs 1b–c). Especially in the male, the caudal fin is markedly lanceolate, and the middle rays may be

prolonged beyond the fin membrane (Fig. 1c). Pseudosphromenus dayi is also slightly more colourful than P.

cupanus, and presents two dark stripes along the side.

Both species are easy to keep and rear in aquaria, and have been in the aquarium hobby since the early 20th

Century. Pseudosphromenus cupanus was imported as an aquarium fish to Germany in 1903 (Rachow, 1936b),

followed by P. dayi in 1907 (Scholze & Pötzschke, 1930) or 1908 (Rachow, 1936a). The latter was then recognized

as being the same as a form of P. cupanus, distinguished but not named by Day (1865:135, 1877:371) (Poenicke,

1908a; Köhler, 1908), and was long treated as a variety or subspecies of P. cupanus (Köhler, 1908; Engmann, 1909;

Rachow, 1936a), only occasionally considered a separate species (e.g., Stansch, 1911; Vierke, 1988). Kottelat

(1994) reviewed the nomenclature of P. dayi, and established its species rank. Although authorship of P. dayi is

usually attributed to Köhler (1908), the name was made available by Engmann (1909) as explained by Kottelat

(1994).

Both species were first described in the genus Polyacanthus Cuvier, a synonym of Trichogaster Schneider, but

contained in Macropodus La Cepède from Regan’s (1909) revision of the Asian anabantoid fishes until Vierke

(1975) resurrected Pseudosphromenus. Pseudosphromenus dayi occurs in Kerala, India (Cherian et al., 2001;

Dahanukar & Rema, Devi 2013; Kottelat, 1994), whereas P. cupanus is found both in Sri Lanka (Pethiyagoda,

KULLANDER ET AL.576 · Zootaxa 3990 (4) © 2015 Magnolia Press

1991), and in southern India (Cherian et al., 2001; Day, 1877:371). The taxonomy of P. cupanus and P. dayi was

addressed most recently by Cherian et al. (2001), comparing a sample of each species. In the phylogenetic analysis

of the Osphronemidae by Rüber et al. (2006), P. cupanus and P. dayi are sister species with an estimated divergence

at 5.3 MYA.

In an assessment of the genetic diversity of the Bangladeshi freshwater-fish fauna we noted that Rahman

(1989, 2005) and Rahman & Ruma (2007) report Pseudosphromenus cupanus from Bangladesh and give also some

fin counts to support the identification. The figure provided in Rahman (1989) is redrawn from the illustration in

Day (1877, pl. 78, fig. 4). The associated images in Rahman (2005) and Rahman & Ruma (2007), however, show

different specimens of Badis Bleeker, probably Badis badis (Hamilton). The latter is reproduced here as Figure 1d.

Further research into the literature on south Asian fishes shows that P. cupanus has been reported with the

distribution Eastern India, Sri Lanka, western Burma, Malay Peninsula and Sumatra by Talwar & Jhingran (1991:

1002–1003), and they are copied by Rahman & Ruma (2007). Pseudosphromenus cupanus is listed in Jayaram

(1981: 382) with the locality information Kerala, Coromandel coasts, Bangladesh, Pakistan, Sri Lanka, Malaya,

and Malay Archipelago. In Jayaram (1999:438–439) the locality information for the genus is copied from that of

Macropodus in Jayaram (1981), i.e., including species of Macropodus in Viet Nam and China and the incorrect

locality in Java for the Chinese species M. opercularis (Linnaeus) in Weber & de Beaufort (1922:347); the

distribution for P. cupanus is expanded with a Maharashtra locality and dropping Malaysia and the Malay

Archipelago. Similar information is repeated in FishBase (Froese & Pauly, 2015) and the IUCN Red List

(Abraham, 2013). Considering that P. cupanus is a species endemic to Sri Lanka and southern India (Cherian et al.,

2001; Day, 1877; Pethiyagoda, 1991) there seems to be a mistake ‘going viral’ here; the objective of this paper is

rectify distribution information for species of Pseudosphromenus and investigate the sources of the confusion.

Material and methods

Distribution data were assembled from the literature and the sources of statements were checked against literature

records and museum collections online in FishNet II and GBIF. Coverage of aquarium literature is selective,

focusing on widely available books and early periodicals. Distribution records in German-language sources were

translated to English. References may use the generic names Polyacanthus or Macropodus for species of

Pseudosphromenus, but we use Pseudosphromenus consistently. Museum codes are: ANSP (Academy of Natural

Sciences of Philadelphia), BMNH (The Natural History Museum, London), MCZ (Museum of Comparative

Zoology, Cambridge), MNHN (Muséum national d’Histoire naturelle, Paris), NRM (Swedish Museum of Natural

History, Stockholm), RMNH (Naturalis Biodiversity Center, Leiden), ZMA (Zoölogisch Museum Amsterdam;

now in RMNH).

The following specimens (*) or database records (**) were assessed:

Badis sp.: ANSP 93953**, 1. Burma: Shinbwiyang. J.W.H. Rehn, 29 May 1945.—ANSP 93954**, 3. Burma:

Shingbwiyang. J.W.H. Rehn, 17 Apr 1945.—ANSP 93959**, 1. Burma: Cahill Creek, J.W.H. Rehn 10 May 1945.

Pseudosphromenus cupanus. NRM 12251*, 1. India, Kerala, Kottayam District, Meenachil River basin, tributary

10 km NW of Kottayam, Olassa village. E. Åhlander, 15 Dec 1987.—NRM 26169*, 22. Sri Lanka, Southern

Province, Point de Galle. O. Nordqvist & A. Stuxberg, 16–21 Dec 1879.—NRM 40344, 7*. Calcutta, Deepak

Nopany's Fish Farm 'Asian Exporter'. F. Fang & A. Roos, 17 Jan 1998.

Pseudosphromenus dayi: BMNH 1909.7.16.35-37**, 3. Malacca. J. P. Arnold.—BMNH 1908.10.26.2-3**, 2.

Malacca. W. Wolterstorff (Syntypes).—BMNH 1933.9.2.8, 1. Sumatra. F. Mayer.—NRM 12069*, 2. India, Kerala,

Kottayam Municipality, Kottayam, Meenachil River 1 km from railway station, riverside. E. Åhlander, 22 Mar

1990. RMNH (ZMA 124.107**), 12. Indonesia, Anak Laut, bergrivierte op Pulo Weh. P. Buitendijk. (Photograph

examined.)

Trichogaster lalius (Hamilton): MCZ 60876**, Pakistan, Sind,Thetta; SE of Karachi. H. Field, 19 Jul 1955.

(Photograph examined.)

Registered as P. dayi: MNHN 1911-0242**, 1. Indo-Malaisie. Lefebvre. (Not further investigated, apparently

aquarium specimen:)

Registered as P. cupanus: MCZ 154985**. 1, West Java. K. Liem, 1955. (The specimen could not be located;

K. Hartel, pers. comm.)

Zootaxa 3990 (4) © 2015 Magnolia Press · 577PSEUDOSPHROMENUS CUPANUS

Results

In the scientific secondary literature the earliest mention of Pseudosphromenus cupanus as occurring in

Bangladesh, Pakistan, Malaysia and Indonesia indeed seems to be that of Jayaram (1981), and the earliest mention

of P. cupanus from Burma seems to be in Talwar & Jhingran (1991), although Burma had been listed in aquarium

literature already by Arnold (1949) and Sterba (1959). The species has not been reported in faunal works on the

fishes of Pakistan (Mirza 1990), Malaysia (Mohsin & Ambak, 1983), or Indonesia (Kottelat et al., 1993), which are

three ichthyologically relatively well-investigated countries. The species is also not listed in the country

monographs for Thailand (Smith, 1945) and Cambodia (Rainboth, 1996). With the exception of the doubtful record

in Weber & de Beaufort (1922), there are no reliable museum records of species of Pseudosphromenus from

outside the native range in southern India and Sri Lanka, and also no literature records of occurrence outside the

native range in India and Sri Lanka except for the unverified records cited here, and which may be phrased in

different ways in secondary references.

The record from Sumatra probably derives from Weber & de Beaufort (1922: 346), who reported material from

Lake Anak laut, on Pulu Weh (= Aneuk Laot, Pulau Weh) near Sumatra. The sample of 12 specimens was

originally deposited in the Zoölogisch Museum Amsterdam (ZMA 124.107), and are now in the Naturalis

Biodiversity Center in Leiden. A photograph of the sample shows it to consist of P. dayi rather than P. cupanus.

The handwritten label states that they were from “Anak Laut, bergrivierte op Pulo Weh”, i.e., from a mountain

stream rather than the lake, and collected by Peter Buitendijk. Buitendijk (1870–1932) was a ship’s surgeon on the

route Amsterdam–Jakarta 1900–1931. He collected fishes and other aquatic animals for the Rijksmuseum van

Natuurlijke Historie in Leiden (now Naturalis Biodiversity Center). Apparently Buitendijk also kept live fishes for

the Amsterdam Zoo (now Natura Artis Magistra) (Fransen et al., 1997), which suggests one source of confusion. It

may be that aquarium stock of P. dayi was confused with some fish collected by Buitendijk on Pulau Weh. Except

for Hoedeman (1954: 477), and derived editions of that book and tertiary references to “Way island” in German

aquarium literature, no subsequent work on anabantoids or Indonesian fishes has cited this record. Hoedeman was

curator at the ZMA. The fin counts provided by Weber & de Beaufort agree with Pseudosphromenus and with no

other Indonesian osphronemid species. It seems highly unlikely that there was or is a natural population of P. dayi

on a small island in the Indonesian archipelago. More likely, the locality information for ZMA 124.107 is in error.

The original reference to occurrence in Myanmar, reported by Talwar & Jhingran (1991), could not be traced,

but there are three samples in the ANSP collection, from Shingbwiyang (= Shin Bway Yang) on the upper

Chindwin River in Myanmar, collected in 1945 by John W.H. Rehn and identified but never published on by Henry

W. Fowler. These specimens were examined for us by Mark Sabaj Pérez at ANSP and identified as a species of

Badis.

The Natural History Museum, London, has samples of Pseudosphromenus dayi labelled Malacca donated by

Walter Wolterstorff in 1908 and Johann Paul Arnold in1909, and also one specimen with locality Sumatra donated

by Fritz Mayer in 1933. Those specimens were donated by aquarists, and the first two are obviously descended

from the first importation of P. dayi. Regan (1909) included those specimens in P. cupanus in his revision of the

Asian anabantoids, where the distribution of P. cupanus is given as “India; Ceylon; Malay Peninsula; in lowland

streams and estuaries”. The Malaysian distribution is certainly attributable to Arnold’s and Wolterstorff’s

specimens. Mayer’s specimen was accessioned along with numerous other aquarium specimens from different

continents (R. Britz, pers. comm.), rendering the locality information open to doubt given that no further records

from Sumatra exist.

The original reference to occurrence in Pakistan in Jayaram (1981) could not be identified but one specimen in

the MCZ collection, collected in Thetta, Pakistan, in 1955, was identified as P. cupanus. An image of the specimen

shows it to be a bleached specimen of Trichogaster lalius. This record seems not to have been observed in the rich

literature on fish biogeography in Pakistan (e.g., Mirza, 2006), and is unlikely to have been the basis for Jayaram’s

record.

An original report on P. cupanus in Bangladesh cannot be found. The species is not mentioned in Bhuiyan’s

(1964) monograph of fishes from Dhaka, one of the earliest accounts of freshwater fishes from Bangladesh. The

earliest local record (Rahman, 1989) is apparently based on the mention in Jayaram (1981).


FIGURE 1 (a) Pseudosphromenus cupanus, NRM 40344, 27.1 mm SL, India, Kolkata, ornamental fish farm; (b) P. dayi, NRM

12069, 22.7 mm SL, India, Kerala, Kottayam; (c) P. dayi, living specimen in aquarium, ca 40 mm SL; (d) Badis badis, image

used as illustration of Pseudosphromenus cupanus in Rahman & Ruma (2007), slightly adjusted. Photo by Gawsia W.

Chowdhury.


The confusion seems to stem to some extent also from aquarium literature. Both P. cupanus and P. dayi have a

long history as aquarium fishes. The three major influential sources of aquarium information were Holly et al.

(1927–1966), Arnold & Ahl (1936), and Sterba (1959). These books had a wide distribution (Sterba was also

translated to English), captured all available information, and were sources for derivative works by others. In the

anglophone domain Axelrod & Schultz (1955) had a similar position.

The origin of the first importation of P. dayi to Germany in 1907 or 1908 was reported initially to be either

Cochin or Malacca (Rachow, 1936b). Although the importer Scholze & Pötzschke had stated that it came from

Cochin (=Kochi, eastern India) (Poenicke, 1908b), Engmann (1909) gave the origin as “Malakka”. Rachow

(1936b) summarized the distribution as “Nearer India: Canara-District (?), Cochinchina, Malay Peninsula and Way

Island (at Sumatra)”. “Way Island” is probably Pulau Weh, and “Nearer India” equals the Indian subcontinent

(including India, Sri Lanka, Pakistan, Nepal, Bhutan, and Bangladesh). Cochinchina equals today’s southern Viet

Nam, and apparently Rachow confuses Cochin on the western coast of India with Cochinchina.

Pseudosphronemus cupanus were imported as aquarium fish to Germany in 1903. Rachow (1936a) gives the

following distribution for P. cupanus: “India, Malabar and Coromandel Coast, Ceylon, allegedly from Siam,

Cochinchina and the Malay Peninsula; mostly in coastal areas and particularly common in small and tiny waters,

also such that are influenced by flood and tide, in irrigation canals of fields and such.”

Arnold & Ahl (1936) describe the distribution of P. cupanus as “Nearer India including Ceylon and Farther

India”, where “Farther India” equals Indochina plus Myanmar. They put P. dayi correctly on the Malabar Coast

(Kochi). Axelrod & Schultz (1955) list Malabar and Coromandel coasts as the native area for P. cupanus, and

Ceylon, Sumatra and Malaya [=Malaysia] as the distribution of P. dayi.

Sterba (1959:616) describes the distribution of P. cupanus cupanus as “Nearer India, Ceylon close to the

coast”; and that of P. cupanus dayi as the “Malabar Coast, Burma, South Viet Nam and Way Island”. The record

from Burma may be from Arnold (1949), the earliest aquarium literature record of P. dayi from Burma that we

could find. Arnold lists “Malabar coast, Burma and Cochinchina” as distribution of P. dayi. In two other editions of

Sterba’s book P. dayi is reported from “Malabar coast, Burma and Cochinchina” (Sterba, 1955:276), and “South

Viet Nam, Way Island” (Sterba, 1987:867).

All in all, aquarium sources report the distribution of P. cupanus as southern India (Western and Eastern

Ghats), Sri Lanka and Indochina; and that of P. dayi as Malabar Coast, Western Ghats, Malaysia, Myanmar and

South Vietnam. We have not found an aquarium literature source for Pakistan or Bangladesh.

Not all secondary and tertiary sources present an erroneous distribution for species of Pseudosphromenus.

Cherian et al. (2001), Kottelat (1994) and Menon (1999) report reliable distributions for both species, except the

latter insisting on Burma for P. dayi. Vierke (1988:114), in a popular work, already rejected the wide distribution of

P. dayi, considering it correctly to occur only in southern India.

Discussion

Pseudosphromenus cupanus is a very distinctive species, and unlikely to be confused with any other osphronemid

species in Bangladesh or Myanmar. Day (1877:371) reports P. cupanus from the Malabar (Kerala) and Coromandel

(Tamil Nadu) coasts. He gives the dorsal and anal fin counts as D. XIV–XVII. 5–7, A. XVI–XIX.9–11 in the

description (XIV–XVI.5–7 and XVI–XIX.10–11, respectively, in the synopsis). With the long dorsal and anal fin it

cannot be confused with Trichopsis vittata, which has a much shorter dorsal fin; or Badis, with only three anal-fin

spines. Since the species data reported by Rahman (1989, 2005), Rahman & Ruma (2007), and Talwar & Jhingran

(1981) are probably taken from literature they are of limited value as evidence of the records from outside Sri

Lanka and southern India. The common osphronemid species in Bangladesh, Trichogaster fasciata (Schneider), T.

chuna (Hamilton), and T. lalius, also with long dorsal and anal fins, are much more deep-bodied than both P.

cupanus and P. dayi. Because there is no species in Bangladesh or Myanmar with which P. cupanus or P. dayi can

be confused, there is no reason to consider that records from those countries are based on misidentified specimens.

The misidentified museum records from Pakistan (MCZ) and Myanmar (ANSP) are an exception, but they were

never published on and consequently did not enter into the literature. The photographs of specimens of Badis in

Rahman (2005) and Rahman & Ruma (2007) are not necessarily to be considered as vouchers. Several of the

species accounts in Siddiqui et al. (2007) are illustrated with photos of some other species. It is unlikely that an


experienced ichthyologist such as Ataur Rahman would have identified a specimen with three anal-fin spines as a

species with 16–19 anal-fin spines.

It may be possible to find additional aquarium references and track the confusion in detail, but, it seems

sufficient here to conclude first that the information in Jayaram’s, Rahman’s, and Talwar & Jhingran’s accounts are

probably derived partly from confused aquarium sources. Both Jayaram (1999) and Talwar & Jhingran (1991) refer

to aquarium literature in comments on the synonymy and generic placement of P. cupanus. One potential mistake

concerns the origin of the first import of P. dayi, Cochin, which should be understood as Kochi on the western

(Malabar) coast of India and which is within the natural range of P. dayi. As evident from Rachow (1936b) it has

been confused with Cochinchina (Viet Nam) where there are no Pseudosphromenus. We have not found authentic

literature sources for Pakistan, Bangladesh or Myanmar. Those records may have been deduced from area

descriptions such as Nearer India and Farther India used in the 19th

and early 20th

Century, although we then miss

references to Bhutan and Nepal. Alternatively, perhaps Myanmar and Bangladesh were added to the list of

countries of occurrence simply to fill the gap between Malaysia and India. Bangladesh was part of Pakistan from

1947 till 1971, but since the species does not occur in either present-day Pakistan or Bangladesh, this name change

cannot have been a source of confusion.

Adding to the problem, all Indian and Bangladeshi sources fail to recognize P. dayi as a distinct species, so the

false distribution data for P. dayi in Sterba (1959) is added to the false distribution data for P. cupanus. To some

extent there may also be some confusion with species of Parosphromenus Bleeker, which look somewhat similar,

but which only occur in Southeast Asia, including Sumatra.

From the case of the garbled distribution data of P. cupanus we learn three things: First, that aquarium

literature is not always reliable, but usually represents compilation from many sources without critical scrutiny.

Aquarium literature is not scientific: it is by default somewhat less reliable in terms of precise facts, and often

based on oral communication where published information is not an alternative. This is not to say it is useless,

because for the most part it is sound and reliable. Nevertheless, as a foundation for national checklists aquarium

literature may be a bad choice. Second, the same applies to popular handbooks and field guides. Third, we cannot

assume anything about distribution without voucher specimens of reliable provenance identified by specialists. In

the case of the Bangladeshi (and Myanmar) Pseudosphromenus, no specimens are on record. We have also not

been able to find evidence of correctly-identified published vouchers for Malaysia, Indochina, Indonesia, or

Pakistan.

A significant aspect of the confusion is also the application of subspecific epithets. Pseudosphromenus

cupanus and P. dayi are distinct taxa with different distribution whether they are considered as subspecies or

species. When the two are synonymized (as done by, e.g., Regan, 1909; Jayaram, 1981; Talwar & Jhingran, 1991;

Rahman, 1989, 2005) they have the combined distribution. This becomes problematic when there are different

resolutions over species status of the concerned taxa, resulting in different concepts of the same name. The P.

cupanus sensu Rahman (1989, 2005) thus represents both P. dayi and P. cupanus, whereas P. cupanus sensu

Pethiyagoda (1991) excludes P. dayi. Thus, following the concept of P. dayi and P. cupanus as different species,

one must consider P. cupanus sensu Rahman as two taxa based on his distribution data, and it becomes unclear

whether it is P. dayi or P. cupanus that does not exist in Bangladesh. It is an illusion of a trinomen that that it can be

abbreviated to just the specific name (the name of the nominotypical subspecies), and therefore recommendable

never to use just the specific name for a polytypic species, but retain the trinomen, or better yet, consider to

abandon the use of trinomina. Trinomina have been next to abandoned from ichthyology following Rosen (1978).

Out of 33032 valid fish species recognized in FishBase (Froese & Pauly, 2015), only 59 species include 2–6

subspecies, many of them obsolete. The use of subspecies is contentious in taxonomy as a whole and has been

contested as being inconsistently applied and/or incongruent with genetic variation and/or lacking evolutionary

context (e.g., Gippoliti & Amori, 2007; Kodandaramaiah et al., 2012; Kullander, 1999; Sangster, 2014; Wilson &

Brown, 1953; Zink, 2004). The case of P. cupanus is further testimony to the confusing nature of trinomina.

To add to the confusion P. cupanus is now reported as extinct in Bangladesh (Hossain, 2014). That is perhaps

the best solution, as it removes the enigma. Several of the species listed as extinct by Hossain (2014), however,

either never occurred in Bangladesh (e.g., Pangio oblonga (Valenciennes)), are misidentified but still very much in

existence (e.g., Danio dangila (Hamilton)), or are not extinct at all (the majority). It is understandable, however,

that a species may be categorized as extinct if it is listed by several authoritative sources, but fails to show up in

inventories. Another recent source, however, lists P. cupanus as common in one site in Bangladesh (Islam, 2012),


but may be based on literature. Pseudosphromenus cupanus is not reported as extinct anywhere else than in

Bangladesh. It is not even threatened anywhere. The information in the global IUCN assessment (Abraham 2013)

is just copied from literature (citing the same wide distribution as Talwar & Jhingran 1991) and is not authoritative.

It is probably correct in the classification as Least Concern, however. If P. cupanus does not exist in Bangladesh it

is not because it has become extinct, but because it never occurred there.

A consequence of erroneous distribution data is that the Bangladeshi list of extinct species now contains one

too many species. However, there is a lesson here that threat status has to be evaluated in a more thorough manner,

searching first for the evidence of the historical existence of the species. Red listing and protection measures cost

time and money which should not be spent on ghost fishes. Consider only the possibility that a major programme

had been started to re-introduce Pseudosphromenus in Bangladesh.

That information about geographical distribution of species of Pseudosphromenus is non-trivial is also

exemplified by the inclusion of P. cupanus in a survey of fish species feeding on mosquito larvae by Chandra et al.

(2008), citing the distribution of P. cupanus as“Eastern India, Sri Lanka, Western Burma (now Myanmar), Malay

Peninsula and Sumatra. Larvivorous fishes may be important agents in biological control of malaria, and as such

introduced in new areas or boosted in areas of natural occurrence (Kamareddine, 2012). Introductions for

biological control may affect natural faunas adversely, and must rely on extremely precise and reliable information.

Use of false distribution data in decisions on harvesting or application could harm the indigenous fauna or be

ineffective for the intended purpose.

We have assumed here that there are no Pseudosphromenus in Bangladesh. We assume so because there is no

evidence that there were ever any. There are no specimens known to be present in local collections, and our own

country-wide surveys in 2014 and 2015 failed to find any. There is also no evidence of this genus in Myanmar or in

South-east Asia. This does not exclude the possibility that there are Pseudosphromenus in the region. Perhaps they

are not reported because they are already in the books (without evidence), or because published illustrations are

misleading, depicting other species. Pseudosphromenus cupanus is or at least has been exported as an ornamental

fish from Kolkata (NRM 40344), relatively close to the Bangladesh/India border, and there is a potential risk of

escapes of the species from holding facilities in Kolkata expanding eastward. There are, however, no records of P.

cupanus from West Bengal (cf. Sen, 1992), the Indian state immediately west of Bangladesh. The genus is absent

from species catalogues for the northern Indian states of Uttar Pradesh and Bihar (Srivastava, 1998) and the

northeastern Indian states (Assam, Arunachal Pradesh, Manipur, Meghalaya, Mizoram, Nagaland, Sikkim, Tripura)

(Vishwanath et al., 2007), several of which border with Bangladesh or Myanmar. Given that there is no recent

revision of the genus covering the entire distribution and including type material, there is a possibility that it may

contain more species diversity, further complicating the issue of which particular species of Pseudosphromenus

does not exist where the genus does not exist. It is, however, even more unlikely that still undiscovered species will

be found within areas of concluded non-existence.

We thus offer illustrations of both P. dayi and P. cupanus here as a standard of reference for local fishwatchers,

as an aid to recognition and discovery of these and potential new species.

Acknowledgements

We are grateful to Ralf Britz (BMNH), Karsten Hartel (MCZ), Martien van Oijen and Ronald de Ruiter (RMNH),

and Mark Sabaj Pérez (ANSP) for information about specimens in their respective collections; Ralf Britz also for

literature; Maurice Kottelat for discussion on the Sumatran locality of P. cupanus; and Gawsia W. Chowdhury for

permission to use her photograph (Fig. 1d). Specimen data from the ANSP and MCZ were accessed through the

Fishnet2 Portal (www.fishnet2.org, 2015-04-25). Specimen data from the BMNH, MNHN, and MCZ were

accessed from the Global Biodiversity Information Facility (GBIF, 2015). The research for this paper was

conducted for the project “Genetic characterization of freshwater fishes in Bangladesh using DNA barcodes”

(Swedish Research Council, contract D0674001 to Sven Kullander and Abdur Rob Mollah).


References

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www.iucnredlist.org (accessed 23 April 2015)

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